Subjective value of risky foods for individual domestic chicks: a hierarchical Bayesian model

For animals to decide which prey to attack, the gain and delay of the food item must be integrated in a value function. However, the subjective value is not obtained by expected profitability when it is accompanied by risk. To estimate the subjective value, we examined choices in a cross-shaped maze with two colored feeders in domestic chicks. When tested by a reversal in food amount or delay, chicks changed choices similarly in both conditions (experiment 1). We therefore examined risk sensitivity for amount and delay (experiment 2) by supplying one feeder with food of fixed profitability and the alternative feeder with high- or low-profitability food at equal probability. Profitability varied in amount (groups 1 and 2 at high and low variance) or in delay (group 3). To find the equilibrium, the amount (groups 1 and 2) or delay (group 3) of the food in the fixed feeder was adjusted in a total of 18 blocks. The Markov chain Monte Carlo method was applied to a hierarchical Bayesian model to estimate the subjective value. Chicks undervalued the variable feeder in group 1 and were indifferent in group 2 but overvalued the variable feeder in group 3 at a population level. Re-examination without the titration procedure (experiment 3) suggested that the subjective value was not absolute for each option. When the delay was varied, the variable option was often given a paradoxically high value depending on fixed alternative. Therefore, the basic assumption of the uniquely determined value function might be questioned.     

A motor-driven feeder for operant training in song birds

Conventional pigeon feeders can make noise, spill food, and injure birds in applications using small song birds as subjects. We developed a motor-driven feeder based on the principle of the piston to address these concerns.     

A pellet feeder for the birds

An alternative means of delivering food to pigeons in operant conditioning research is described. The feeder allows greater control of the amount of food delivered and reduces the amount of time necessary for the pigeon to collect the food. It is possible to extend the length of experimental sessions due to the reduction of food intake. Data obtained using the pellet feeder indicated that the control of responding is comparable to that observed with the standard grain-magazine feeder.     

Pigeons discriminate between human feeders

Considered as plague in many cities, pigeons in urban areas live close to human activities and exploit this proximity to find food which is often directly delivered by people. In this study, we explored the capacity of feral pigeons to take advantage of this human-based food resource and discriminate between friendly and hostile people. Our study was conducted in an urban park. Pigeons were fed by two experimenters of approximately the same age and skin colour but wearing coats of different colours. During the training sessions, the two human feeders displayed different attitudes: one of the feeders was neutral and the second was hostile and chased away the pigeons. During the two test phases subsequent to the training phase, both feeders became neutral. Two experiments were conducted, one with one male and one female feeder and the second with two female feeders. In both experiments, the pigeons learned to quickly (six to nine sessions) discriminate between the feeders and maintained this discrimination during the test phases. The pigeons avoided the hostile feeder even when the two feeders exchanged their coats, suggesting that they used stable individual characteristics to differentiate between the experimenter feeders. Thus, pigeons are able to learn quickly from their interactions with human feeders and use this knowledge to maximize the profitability of the urban environment. This study provides the first experimental evidence in feral pigeons for this level of human discrimination.     

Choice as a dependent measure in autoshaping: sensitivity to frequency and duration of food presentation.

Previous investigations have shown that rate, latency, and percentage of trials with at least one response are somewhat insensitive measures of the strength of autoshaped responding. In the present studies, these measures were contrasted with the allocation of responding during simultaneous choice tests, a measure of response strength frequently used in operant paradigms. In two experiments, nine pigeons were exposed to a forward pairing autoshaping procedure. Training sessions consisted of the successive presentation of three stimuli, each followed by food on either 100%, 50%, or 0% of the trials. Choice testing involved the simultaneous presentation of the three stimuli. In Experiment I, all pigeons consistently directed their initial choice responses and the majority of subsequent responses to the stimulus always followed by food, despite the fact that during training sessions the response rates of most birds were highest in the presence of the stimulus followed by food on 50% of the trials. In Experiment II, rate, latency, and percentage of trials with at least one response did not change appreciably as a function of duration of feeder presentations. However, choice responding was lawfully affected by duration of feeder presentations. These data suggest that choice is perhaps a more sensitive measure of the strength of autoshaped responding than other, more commonly employed, indices.     

Two-key concurrent responding: response-reinforcement dependencies and blackouts

Two-key concurrent responding was maintained for three pigeons by a single variable-interval 1-minute schedule of reinforcement in conjunction with a random number generator that assigned feeder operations between keys with equal probability. The duration of blackouts was varied between keys when each response initiated a blackout, and grain arranged by the variable-interval schedule was automatically presented after a blackout (Exp. I). In Exp. II every key peck, except for those that produced grain, initiated a blackout, and grain was dependent upon a response following a blackout. For each pigeon in Exp. I and for one pigeon in Exp. II, the relative frequency of responding on a key approximated, i.e., matched, the relative reciprocal of the duration of the blackout interval on that key. In a third experiment, blackouts scheduled on a variable-interval were of equal duration on the two keys. For one key, grain automatically followed each blackout; for the other key, grain was dependent upon a response and never followed a blackout. The relative frequency of responding on the former key, i.e., the delay key, better approximated the negative exponential function obtained by Chung (1965) than the matching function predicted by Chung and Herrnstein (1967).     

Duration discrimination is better with food access as the signal than with light as the signal

In Experiment 1 a go/no-go discrimination procedure was used to compare control of five pigeons' keypecking by food-access duration with control by light duration. Pecks to an illuminated key were reinforced with grain following 10-sec presentations of food access or houselight, but not after 5-sec presentations of either stimulus. Each subject discriminated food-access duration faster and to a greater degree than light duration. In four between-subject replications, pigeons discriminated food-access duration better than the duration of a localized light, the feeder light and a keylight, and with either water or food as reinforcement. In Experiment 2 control by durations of food access and light was compared using a conditional right-left choice procedure (two pigeons), and a delayed symbolic matching-to-sample procedure (six pigeons). Under both, choice accuracy again was higher on food-access trials. The results of Experiment 3, in which two pigeons received generalization trials with durations of food access and light that were intermediate to the training values, confirmed that responding was controlled by the duration dimension of both food access and light. The superior control by food access is consistent with previous evidence that food is an effective and memorable stimulus, possibly because of its biological importance. These results also provided empirical support for the commonly made assumption that stimuli differ in effectiveness. As well, the results show that the stimulus to be discriminated can play an important role in the accuracy of duration disciminations, a fact which has implications for the study of temporal discriminations in animals.     

Incentive theory: IV. Magnitude of reward

Incentive theory is successfully applied to data from experiments in which the amount of food reward is varied. This is accomplished by assuming that incentive value is a negatively accelerated function of reward duration. The interaction of the magnitude of a reward with its delay is confirmed, and the causes and implications of this interaction are discussed.     

Tits use amodal completion in predator recognition: a field experiment

Amodal completion enables an animal to perceive partly concealed objects as an entirety, and to interact with them appropriately. Several studies, based upon either operant conditioning or filial imprinting techniques, have shown that various animals (both mammals and birds) can perform amodal completion. Before this study, the use of amodal completion by untrained animals in the recognition of objects had not been considered. Using two feeders, we observed in a field experiment the reaction of tits to the torso of a sparrowhawk (partly occluded or an ‘amputated’ dummy) in two different treatments (sparrowhawk torso vs. complete dummy pigeon; and torso vs. complete dummy sparrowhawk). It is clear that the birds considered the two torso variants as predators and kept away from both of them when the second feeder offered a ‘pigeon’ instead. On the other hand, when a ‘complete sparrowhawk’ was present on the second feeder, the number of visits to the occluded torso remained low; while the number of visits to the amputated one increased threefold. Birds risked perching near what was clearly an amputated torso; while the fear of a “hiding” (occluded) torso remained unchanged, when the second feeder did not provide a safe alternative. Such discrimination between torsos requires the ability for amodal completion. Our results demonstrate that in their recognition process, the birds not only use simple sign stimuli, but also complex cognitive functions.     

Social learning of optimal responding and producer-scrounger strategies in flocks of pigeons

Social learning of optimal responding and producer-scrounger strategies in flocks of pigeons. This experiment evaluated the effect of varying the effort required in order to access food upon the acquisition of a novel response, and upon utilizing producer-scrounger strategies, in flocks of pigeons exposed to an expert conspecific. Flocks of na?ve observers were exposed to a pigeon trained to open the seals on food deposits. The colors of the seals were correlated with differing required efforts. Later, the trained pigeon was removed while the color-effort correlation was maintained. The results showed that the observers acquired the response of opening the seals and that they executed it discriminatingly, responding in a greater proportion to the color that signaled the least required effort. They also assumed the roles of both producers and scroungers, which varied as a function of the composition of the group. The data obtained are congruent with prior findings regarding social learning, and also with the concepts proposed by the optimal foraging theory.     

Testing a stochastic foraging model in an operant simulation: Agreement with qualitative but not quantitative predictions

An operant simulation of foraging through baited and empty patches was studied with 4 pigeons. On a three-key panel, side keys were designated as patches, and successive opportunities to complete 16 fixed-ratio 10 schedules on side keys were defined as encounters with feeders. In a random half of the patches in any session, some of the fixed-ratio 10 schedules yielded reinforcement (baited feeders) and the other schedules yielded nonreinforcement (empty feeders). In the other half of the patches, all feeders were empty. Pigeons could travel between patches at any time by completing a fixed-ratio schedule on the center key. An optimal foraging model was tested in Experiments 1 and 2 by varying center-key travel time and number of baited feeders in baited patches. The ordinal predictions that number of feeders visited in empty patches would increase with travel time and decrease as number of baited feeders increased were supported, but pigeons visited far more feeders in empty patches than the optimal number predicted by the model to maximize energy/time. In Experiment 3, evidence was found to suggest that the number of empty feeders encountered before the first baited feeder in baited patches is an important factor controlling leaving empty patches.     

Extinction-induced aggression

Pigeons were conditioned to peck a response key under a procedure that alternated periods of food reinforcement with periods of extinction. The pigeons attacked a nearby pigeon at the onset of extinction. Some also attacked a stuffed model of a pigeon. The duration of attack was an inverse function of the time since the last food reinforcement and a direct function of the number of reinforcements. The pigeons attacked after the last food delivery whether or not the conditioned pecking response was required and whether or not the extinction period was signaled. The food had to be eaten; the mere sight and sound of food being delivered did not produce attack. Prior satiation reduced attack. The phenomenon was not attributable to a past history of competition between pigeons since socially deprived pigeons also attacked. Superstitious reinforcement of attack was not found to be a factor. The results indicated that the transition from food reinforcement to extinction was an aversive event that produced aggression.     

Delay discounting in the pigeon: In search of a magnitude effect

The magnitude effect, where larger outcomes are discounted proportionally less than smaller outcomes, is a well‐established phenomenon in delay discounting by human participants. To this point in the literature magnitude effects have not been reliably evidenced in nonhuman animals. , however, used a concurrent‐chains arrangement with pigeon and found evidence for a magnitude effect. Grace et al. suggested that in many delay discounting experimental arrangements with nonhuman animals (e.g., adjusting amount, adjusting delay) the organism is not given the opportunity to directly compare outcomes of different sizes. They suggest that because of the lack of direct comparison it is difficult for the organism to determine the relative size of each outcome, which in turn mutes the effect of the amount differences between outcomes. As a test of this “comparison hypothesis,” the present experiment was conducted to assess whether the magnitude effect would be evidenced in pigeon when using an adjusting amount procedure where outcomes of different amounts were presented proximally. In the present arrangement, pigeons were presented two choice panels in an operant chamber where each panel was associated with an independent adjusting amount delay discounting task, but with differing outcome amounts (i.e., a 32‐food pellet panel and an 8‐food pellet panel). In this arrangement the choice panels alternated in their availability within a session from trial block to trial block. The present findings indicate no reliable effect of amount, even when the outcomes were proximal and thus readily comparable. This result suggests that the lack of magnitude effect is not driven by the organism's ability to compare the difference in amount between choice alternatives.     

Response rate, latency, and resistance to change

Pigeons were trained on a multiple variable-interval/variable-interval schedule with pacing contingencies that generated high response rates in one component and low response rates in the other. Timeout periods separated the schedule components. During resistance-to-change tests, response-independent food was presented during the timeout periods, and the duration of that food presentation was varied among test sessions. Response rates in the schedule components decreased and latencies to the first response increased as a function of the duration of food presentations during the timeout. Both dependent measures changed about the same amount relative to their own baseline levels. The conclusions are that baseline response rates controlled by pacing contingencies are equally resistant to change, given equal reinforcement densities, and latency is a sensitive measure of resistance to change.     

Conditioned suppression/avoidance as a procedure for testing hearing in birds: The domestic pigeon (Columba livia)

Although the domestic pigeon is commonly used in learning experiments, it is a notoriously difficult subject in auditory psychophysical experiments, even those in which it need only respond when it detects a sound. This is because pigeons tend to respond in the absence of sound—that is, they have a high false-positive rate—which makes it difficult to determine a pigeon’s audiogram. However, false positives are easily controlled in the method of conditioned suppression/avoidance, in which a pigeon is trained to peck a key to obtain food and to stop pecking whenever it detects a sound that signals impending electric shock. Here, we describe how to determine psychophysical thresholds in pigeons using a method of conditioned suppression in which avoidable shock is delivered through a bead chain wrapped around the base of a pigeon’s wings. The resulting audiogram spans the range from 2 to 8000 Hz; it falls approximately in the middle of the distribution of previous pigeon audiograms and supports the finding of Kreithen and Quine (Journal of Comparative Physiology 129:1–4, 1979) that pigeons hear infrasound.     

Temporal discrimination and a free-operant psychophysical procedure.

Pigeons were presented a series of keylight time periods (separated by blackouts) during which two response keys were lit, one by blue light and the other either by orange or green. Blue-key responses changed the color on the other key. Orange-key responses sometimes produced food during the first half of a time period; green-key responses sometimes produced food during the second half. In three experiments, the probability of a green-key response increased as a function of elapsed time. Experiment 1 compared performance when the duration of the keylight periods was varied across a wide range. Discrimination of performance was similar across the range of durations. Experiment 2 varied both relative reinforcement rate and the local reinforcement rate for orange-key and green-key responses. These manipulations produced changes in response bias but not discrimination sensitivity. Experiment 3 varied the local temporal placement of reinforcers within time periods and demonstrated that choice behavior was affected by differential reinforcement at different points during the time periods. The results were consistent with previous research on duration discrimination that used psychophysical trials procedures.     

Conditioning of the aggressive behavior of pigeons by a fixed-interval schedule of reinforcement

Operant reinforcement of aggression was studied in food-deprived pigeons by delivering food for attacks against a target pigeon. The food was delivered according to a fixed-interval schedule and attack behavior was recorded automatically. Attack could be conditioned and extinguished, and the proportion of time spent in attack was a direct function of the frequency of reinforcement of the attack. The fixed-interval schedule produced an increasing rate of attack during the interval between food reinforcements. This positive curvature was an inverse function of the duration of the interval. The findings revealed that the duration and temporal patterning of the complex social behavior of attack can be influenced in a substantial and predictable manner by the schedule and frequency of operant reinforcement.     

Effects of search cost on foraging and feeding: a three-component chain analysis

An experiment determined whether pigeons minimize number of key pecks per food delivery and maintain their baseline intake of food while key pecking on a three-component chain schedule. Pigeons at either 80% or 100% body weight obtained all their food during baseline and contingency sessions. During baseline sessions, pecks on the left and center keys had no consequences; each peck on the right key activated the feeder. During contingency sessions, pigeons key pecked on a three-component chain schedule simulating components of a foraging chain. In the search component either 3, 9 or 15 key pecks (varied parametrically across blocks of sessions) on the left key produced a stimulus on the middle key, indicating an encounter with either the low-cost prey (3 key pecks) or an equally probable high-cost prey (21 key pecks). In the procurement component the pigeon pecked either: (a) the left key once, thus returning to the search component, or (b) the middle key either 3 or 21 times, which activated the right response key. In the handling component one peck on the right key operated the feeder. The pigeons always procured the low-cost prey and minimized the number of key pecks per hopper by procuring the high-cost prey when the search-cost ratio was high (15 key pecks) but not when it was low (3 key pecks). All pigeons maintained their baselines of eating during contingency sessions by key pecking more frequently and eating more efficiently. The 80% body-weight birds produced higher overall rates of key pecking and eating. These results have implications for ecological theories of optimal foraging and for psychological theories of learned performance.     

Flexible cue use in food-caching birds

An animal’s memory may be limited in capacity, which may result in competition among available memory cues. If such competition exists, natural selection may favor prioritization of different memory cues based on cue reliability and on associated differences in the environment and life history. Food-caching birds store numerous food items and appear to rely on memory to retrieve caches. Previous studies suggested that caching species should always prioritize spatial cues over non-spatial cues when both are available, because non-spatial cues may be unreliable in a changing environment; however, it remains unclear whether non-spatial cues should always be ignored when spatial cues are available. We tested whether mountain chickadees (Poecile gambeli), a food-caching species, prioritize memory for spatial cues over color cues when relocating previously found food in an associative learning task. In training trials, birds were exposed to food in a feeder where both spatial location and color were associated. During subsequent unrewarded test trials, color was dissociated from spatial location. Chickadees showed a significant pattern of inspecting feeders associated with correct color first, prior to visiting correct spatial locations. Our findings argue against the hypothesis that the memory of spatial cues should always take priority over any non-spatial cues, including color cues, in food-caching species, because in our experiment mountain chickadees chose color over spatial cues. Our results thus suggest that caching species may be more flexible in cue use than previously thought, possibly dependent upon the environment and complexity of available cues.     

A response duration schedule: effects of training, extinction, and deprivation

Rats were trained to hold down a lever for at least 40 consecutive seconds. When the lever had been held down for 40 sec, white noise came on. Releasing the bar in the presence of the noise turned off the noise and operated a feeder that delivered a pellet of food. At the end of training, frequency distributions of response durations peaked at 40 to 41 sec. If as in training, holding down the lever produced white noise at the end of 40 sec, and release of the lever terminated the noise and operated the feeder, but no food delivery occurred, duration distributions and several other measures were initially not very different from when food was delivered. However, if during extinction white noise was never produced by lever holding, and feeder operation did not occur upon lever release, most responses were shorter than 1 sec in duration, some were much longer than 41 sec, and duration distributions did not peak at 40 to 41 sec. When reinforcement was reinstated after extinction, performance quickly returned to pre-extinction measures. Further sessions at different levels of deprivation produced only temporary disruptions in performance.